Comparative Anatomical Studies on Some Species of the Genus Amaranthus (Family: Amaranthaceae) for the Development of an Identiﬁcation Guide Q ⇑ Abbas A

Annals of Agricultural Science xxx (2017) xxx–xxx

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Comparative anatomical studies on some species of the genus Amaranthus (Family: Amaranthaceae) for the development of an identiﬁcation guide q ⇑ Abbas A. El-Ghamery a, Ahmed M. Sadek a, , Ola H. Abd El Bar b a Botany and Microbiology Department, Faculty of Science, Al-Azhar University, Cairo, Egypt b Agriculturale Botany Department, Faculty of Agriculture, Ain-Shams University, Cairo, Egypt article info abstract

Article history: A study of anatomical features of mature leaves and stems (at fruiting stage) of 12 Amaranthus taxa Received 20 September 2016 (Family: Amaranthaceae) shows high variation between them and supplied new characters. The internal Received in revised form 2 November 2016 structures were evaluated to clarify their effectiveness in solving taxonomic complexity and identifica- Accepted 16 November 2016 tion difficulty in this genus. Observation of the transections of blades showed that the epidermis is unis- Available online xxxx eriate, ground tissue consists of angular collenchyma and thin parenchyma. The vascular bundles shape has three patterns crescent, ring, ovate. Also they may be united or separated while the midrib shape in Keywords: cross section has two patterns in which U-shaped, cordate or crescent bundle occurs. All leaves are peti- Amaranthus olate. The examination of the petioles exhibits new and varied characters such as petiole shape (cross sec- Leaf and stem anatomy DELTA key tion), vascular bundles (shape, number, arrangement). While the resulted characters from the Identification observation of the stem structure showed less variation. Nineteen qualitative characters with 38 character states resulted from leaf anatomy. Only (8) characters were sufficient to generate an identification anatomical key. DELTA program was used in key-generation. Also different measurements were carried out by a photo analysis program (Image J), such as lamina thickness, mesophyll thickness, area of upper and lower epidermal cells and thickness of upper and lower epidermal cells to exhibit most possible dis- similarities between the studied species. Ó 2017 Production and hosting by Elsevier B.V. on behalf of Faculty of Agriculture, Ain Shams University. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc- nd/4.0/).

Introduction misapplication of names create intricate problems in its taxonomy (Costea et al., 2001; Iamonico, 2014a, 2014b, 2016a, 2016b; Nestor, The number of Amaranthus species varies in literature. The 2015). genus Amaranthus L. (Amaranthaceae Juss.) consists about 65–80 Furthermore, some Amaranthus species are aggressive annual (Bojian et al., 2003; Mosyakin and Robertson, 1996; Mujica and weeds in the world, causing impacts on the natural ﬂoras and veg- Jacobsen, 2003; Iamonico, 2014a, 2014b.). 455 species names for etation and on cultivated ﬁeld Amaranthus presence can lower the genus Amaranthus are known with the following statistics: grain yield and quality as well as hinder mechanical harvest 105 (23.1%) accepted, 262 (57.6%) synonyms and 88 (19.3%) unas- (Wax, 1995). Some of this species also have been shown to possess sessed (The plant list, 2013). However due to the few studies on allelopathic chemicals that reduce seedling vigor of several crops Amaranthus systematics the number still tentative. Also the and weed species (Menges, 1987, 1988), while others may induce nomenclature of Amaranthus is critical since hundred names were toxicosis and death in dairy cattle (Kerr and Keleh, 1998). The grain published during the centuries and the nomenclatural disorder/ Amaranths are ancient crops with increasing prospects as potential food and feed resources because of their high grain protein and starch quality and high-nitrogen, highly digestible vegetative tis- Peer review under responsibility of Faculty of Agriculture, Ain-Shams University. ⇑ sues (Cai et al., 1998a,b). Some Amaranthus species have medicinal Corresponding author. importance as A. spinosus (Pal et al., 2013). E-mail addresses: [email protected], [email protected] (O.H. Abd El Bar). http://dx.doi.org/10.1016/j.aoas.2016.11.001 0570-1783/Ó 2017 Production and hosting by Elsevier B.V. on behalf of Faculty of Agriculture, Ain Shams University. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

Among the morphological studies on different Amaranthus spe- were immediately fixed in (FAA) according to Johasen (1940). cies; Abdul Rahaman and Oladele (2003) and Gaafar et al. (2015) The third node and internode were selected for the stem study. studied the stomata; Alege and Daudu (2014), Pal et al. (2013), Serial transverse sections (10–20 lm in thickness) were cut with Munir et al. (2011), Hong et al. (2005) studied the leaves. a steel blade on an IHC World KD-1508A rotary microtome and Kraehmer (2013), Carlquist (2003) and Metcalfe & Chalk (1950, fixed on slides by means of Haupt’s adhesive. The sections were 1979) stained with a safranin-fastgreen combination, and then mounted As a consequence, some issues of the taxonomy of Amaranthus in Canada balsam (Sass, 1961). still remain not completely solved, although some approaches Observation and photomicrographs were achieved using XSZ- was carried out (Hauptli and Jain, 1978; Joshi and Rana, 1991; N107 Research Microscope fitted with Premiere MA88-900 digital Bojian et al., 2003, Mosyakin and Robertson, 2003; Hussein, camera. 14 measurements for leaf and stem internal structures, 2005; Pinto and Velasquez, 2010; Iamonico, 2012; Iamonico and epidermal cell area were carried out by the ImageJ Ver. 1.49o pro- Das, 2014). gram (standard software in scientific image analysis) and cali- Because of the poor published data about the leaf and stem ana- brated by using a standard stage micrometer slide. Each recorded tomies, especially concerning the taxonomical implications, we measurement is the average of ten measurements. here present a study aims to evaluate the taxonomic value of leaf Also stomatal density and stomatal index where calculated as and stem structures of some Amaranthus species, with the final described by Wilkinson (1979). aim to facilitate identification Amaranthus and improve the taxonomy of the genus. SI ¼ S 100=S þ E

Various data obtained from the description of the leaf anatomy Material and methods were subjected to automated key generation using version 4.12 of the DELTA suite of program (Dallwitz et al., 2000). Seeds of 11 Amaranthus accessions were supplied by the Royal Botanic Garden at Kew (London, UK) while one taxon (A. tricolor) was selected from one of the Botanical Gardens in Egypt Results (National Garden in Nasr City). 12 species of Amaranthus were studied in the present study (Table 1). All the plants were grown The observation of the internal structure of the leaf and stem of from seeds. the studied species revealed (14) qualitative characters (Table 3). The leaf structure was examined using ﬁve segments (lengths of 28 qualitative characters were resulted and summarized in 1–2 mm) of the blade and petiole (fruiting stage) and then they (Table 2). Also they comprehensively listed in the following text:

Table 1 Species and accessions of Amaranthus used in this study.

No. Species Accession no. Origin Subgenus Habitat 1 A. albus L. 0022453 Germany Albersia Wild 2 A. blitum L. 0006563 France Albersia Cultivated 3 A. caudatus L. 0003919 Unknown Amaranthus Wild 4 A. deﬂexus L. 0020275 Italy Albersia Cultivated 5 A. dubius Mart.ex Thell. 0006574 Colombia Albersia Wild 6 A. graecizans L. 0020242 Portugal Amaranthus Wild 7 A. hybridus L. 0019657 Bolivia Amaranthus Wild 8 A. powellii S. Watson. 0030999 France Amaranthus Wild 9 A. retroﬂexus L. 0008394 Italy Amaranthus Wild 10 A. spinosus L. 0017882 Unknown Amaranthus Wild 11 A. tricolor L. Unknown Egypt Amaranthus Cultivated 12 A. x ozanonii Piszter. 0008383 Unknown Amaranthus Cultivated

Table 2 Qualitative characters of leaf and stem of the studied species of Amaranthus. Character number is preceded by # and states of the same character are assigned serial numbers.

Characters Species A. A. A. A. A. A. A. A. A. A. A. A. x albus blitum caudatus deﬂexus dubius graecizans hybridus powellii retroﬂexus spinosus tricolor ozanonii #1. Petiole shape in TS/1.crescent/2. 11 4 1 1 1 1 1 1 2 2 3 rounded/3.cordate/4.V-shaped #2. Protuberances/1.present/2.absent 1 1 1 1 1 1 1 1 1 1 2 1 #3. Epidermis type/1.uniseriat with 11 1 1 1 1 1 1 1 1 1 1 radial or pueblos cells #4. Ground tissue type/ 11 1 1 1 1 1 1 1 1 1 1 1.parenchymatous and angular collenchyma #5. Sand crystals in petiole/1.present/2. 11 1 1 1 1 1 1 1 1 1 1 absente #6. Number of vascular bundles/1.8/ 1 2 3 4 3 5 4 6 1 7 6 7

Table 2 (continued)

Characters Species A. A. A. A. A. A. A. A. A. A. A. A. x albus blitum caudatus deflexus dubius graecizans hybridus powellii retroflexus spinosus tricolor ozanonii 2.4/3.13/4.6/5.12/6.11/7.7 #7. Vascular bundles arrangement/1. 11 1 3 3 3 3 1 1 1 2 3 crescent/2.circular/3.arc #8. Bundle sheath/1.present/2.absent 1 2 2 1 2 1 1 1 1 1 2 2 #9. Midrib shape in TS/1.crescent/2. 22 2 2 2 1 2 2 1 1 2 1 rounded #10. Ground tissue type/1.parenchyma 11 1 1 1 1 1 1 1 1 1 1 and collenchyma #11. Sand crystals/1.presente/2.absente 1 1 1 1 1 1 1 1 1 1 1 1 #12. Number of Vascular bundles/1.5/ 12 3 1 1 2 4 2 5 6 4 1 2.3/3.9/4.6/5.4/6.2 #13. Vascular bundles shape/1. 13 1 1 2 1 1 1 1 1 2 2 crescent/2.ring/3.ovate #14. Single phloem strand in midrib/1. 21 2 2 2 2 2 2 2 2 2 2 present/2.absent #15. Mesophyll type/1.dorsiventral/2. 11 1 1 1 1 1 1 1 1 2 1 isolaterl #16. Kranz/1.presente/2.absente 1 1 1 1 1 1 1 1 1 1 1 1 #17. Druses/1.presente in wings and 11 1 1 1 1 1 1 1 1 1 1 absent from midrib #18. Trichomes type/1. Uniseriat, 11 1 1 1 1 1 1 2 1 1 1 glandular, multicellular, un-branche/ 2.uniseriat and biseriat, multicellular, glandular, unbranched #19. Type of stomata/1.anomocytic 11 1 1 1 1 2 1 1 1 1 1 only/2.anomocytic and hemiparacytic #20. Shape of internode TS/1.irregular 11 1 1 1 1 1 1 1 1 1 1 ovate/ #21. Epidermis type/1.uniseriate with 11 1 1 1 1 1 1 1 1 1 1 radial cells #22. Cortex tissues/1.collenchyma and 11 1 1 1 1 1 1 1 1 1 1 parenchyma #23. Sand crystals in cortex/1.present 1 1 1 1 1 1 1 1 1 1 1 1 #24. Pith tissue/1.parenchymatous 1 1 1 1 1 1 1 1 1 1 1 1 #25. Anomalous cambial ring around 11 1 1 1 1 1 1 1 1 1 1 the vascular cylinder/1.present #26. Conjugative tissue in vascular 11 1 2 1 1 1 1 1 1 1 1 cylinder/1.thin parenchyma only/2. thin and ligni-fied parenchyma #27. Node type/1.unilacunar with 4 leaf 12 3 2 4 2 2 1 5 1 1 1 traces/2.unilacunar with 3 leaf traces/3. unilacunar with 7 leaf traces/4.unilacunar with 9 leaf traces/5.unilacunar with 5 leaf traces #28. Trichomes type on stem/1. 11 1 1 1 1 1 1 1 1 1 1 Uniseriat, glandular, multicellular, un-branched

Petiole: the petiole shape in cross section is crescent in spe- Lamina: Midrib shape in T.S. crescent in species (6, 9, 10, & 12) cies (1, 2, 4, 5, 6, 7, 8 & 9), rounded in species no. (10 & 11), cor- and rounded in species (1, 2, 3, 4, 5, 7, 8, & 11). In all species ground date in species (12) and V-shape in species (3). Protuberances tissue is parenchyma and collenchyma tissue. Sand crystals present in all species and absent in species (11). In all species present. Number of Vascular bundles 5 in species (1, 4, 5 & 12), 3 epidermis simple, ground tissue parenchymatous and angular in species (2, 6 & 8), 9 in species (3), 6 in species (7 &11), 4 in spe- collenchyma and sand crystals in petiole present. Number of vas- cies (9) and 2 in species (10). Vascular bundles shape crescent in cular bundles 8 in species (1 & 9), 4 in species (2), 13 in species species (1, 3, 4, 6, 7, 8, 9 & 10), ring in species (5, 11 & 12) and ovate (3 & 5), 6 in species (4 & 7), 12 in species (6), 11 in species (8 & in species (2). Single phloem strand in midrib present in species (2) 11) and 7 in species (10 & 12). Vascular bundles arrangement and absent others. Mesophyll type isolaterl in species (11) and crescent in species (1, 2, 3, 8, 9 & 10), circular in species (11) dorsiventral in others. Kranz and druses present in all species and arc in species (4, 5, 6, 7 & 12). Bundle sheath present in spe- (Figs. 5–10). cies (1, 4, 6, 7, 8, 9 & 10) and absent from species (2, 3, 5, 11 & Trichomes type uniseriate, glandular, multicellular, un-branche 12) (Figs. 1–4). in all species except in species (9) it is uniseriat and biseriat, mul-

Please cite this article in press as: El-Ghamery, A.A., et al. Comparative anatomical studies on some species of the genus Amaranthus (Family: Amaran- thaceae) for the development of an identification guide. Ann. Agric. Sci. (2017), http://dx.doi.org/10.1016/j.aoas.2016.11.001 4 A.A. El-Ghamery et al. / Annals of Agricultural Science xxx (2017) xxx–xxx ticellular, glandular, and un-branched. Type of stomata anomocytic species (5) and unilacunar with 5 leaf traces in species (9). Tri- only in all species except species (7) it is anomocytic and hemi- chomes type on stem uniseriate in all species (Plate 1). paracytic. Druses present in the wings and absent from midrib in Eight characters of leaf anatomy were used to generate an all species (Figs. 11–16). anatomical identification key through the DELTA key-generating program and only seven characters were sufficient to produce the key as following:

Key based on anatomical characters of the leaf Characters: 8 indata, 8 included, 7 in key Items: 12 indata, 12 included, 12 in key Characters included: 1–8 Character reliabilities: 1–8,5

1. Number of vascular bundles in petiole 8 2 Number of vascular bundles in petiole 4 Amaranthus blitum Number of vascular bundles in petiole 13 3 Number of vascular bundles in petiole 6 4 Number of vascular bundles in petiole 12 Amaranthus graecizans Number of vascular bundles in petiole 11 5 Number of vascular bundles in petiole 7 6

2(1). Trichomes type multicellular, uniseriate only, glandular and unbranched Amaranthus albus Trichomes type multicellular, uniseriate and biseriate glandular and unbranched Amaranthus retroﬂexus

3(1). Petiole shape in cross section crescent (with two small protuberances); Shape of vascular bundles in Amaranthus petiole crescent dubius Petiole shape in cross section cordate (with two small protuberances); Shape of vascular bundles in Amaranthus petiole V-shaped caudatus

4(1). Types of stomata at the upper epidermis anomocytic only Amaranthus deﬂexus Types of stomata at the upper epidermis anomocytic and hemiparacytic Amaranthus hybridus

5(1). Petiole shape in cross section crescent (with two small protuberances); Mesophyll organization Amaranthus dorsiventral; Shape of vascular bundles in petiole crescent; Bundle sheath in petiole present powellii Petiole shape in cross section V-shaped (without protuberances); Mesophyll organization isolateral; Amaranthus Shape of vascular bundles in petiole U-shaped; Bundle sheath in petiole absent tricolor

6(1). Petiole shape in cross section V-shaped (without protuberances); Bundle sheath in petiole present Amaranthus spinosus Petiole shape in cross section cordate (with two small protuberances); Bundle sheath in petiole absent Amaranthus x ozanonii

Stem Discussion

Internode shape in T.S. of all species irregular ovate, epidermis The results obtained from the present study, especially simple, cortex tissues collenchyma and parenchyma, sand crystals comparison with the previous one reveals that the anatomy of present in cortex, pith parenchymatous, anomalous cambial ring the leaves of the current species can be used to identify some around the vascular cylinder present. Conjugative tissue in vascu- taxa. lar cylinder thin parenchyma only in all species except species (4) The investigators on the characters of stem of some Amaran- it is thin and ligniﬁed parenchyma (Figs. 17 & 18). thus species mostly focused on the internode structure rather Node type unilacunar with 4 leaf traces in (1, 8, 10, 11 & 12), than the nodal structure and also they examined the kranz in unilacunar with 3 leaf traces in species (2, 4, 6 & 7), unilacunar the lamina and relatively neglected the petiole characters. All with 7 leaf traces in species (3), unilacunar with 9 leaf traces in characters exhibited from stem (node and internode) are com-

patible with those reported by Metcalfe and Chalk (1950), Kraehmer (2013), Gaafar et al. (2015), Carlquist (2003) and Costea et al. (2001). The stem showed less variation between m) l

( the studied species. All resulted characters from lamina study tissue

thickness of A. albus, A. caudatus, A. deﬂexus, A. dubius, A. gracecizans, Collenchyma A. hybridus, A. powllii, A. retroﬂexus, A. spinosus, A. tricolor and A. x ozanonii are identical to those reported by Gaafar et al. m) Stem

l (2015), Alege and Daudu (2014), Hong et al. (2005), Muhaidat ( Cortex

thickness et al. (2007), Omosun et al. (2008), Omamni (2005), Tardif et al. (2006), Ferreira et al. (2003), Fisher and Evert (1982) and Baral (2013). The resulted characters from petiole study of A. retroflexus were suitable to those reported by Fisher and (mm) Thired diameter internode Evert (1982), while no researches on the other species were found (see Table 4). Also the resulted anatomical key based on leaf anatomy can be used as a confirmatory key that ensure the identification of lower epidermis %Stomatal the studied taxa and surmount the similarity problem between index of the the taxa of the genus Amaranthus. The software package DELTA is a sophisticated and powerful data-basing program which

2 stores morphological data for export in a number of different forms. In this way it acts as a manager of taxonomic research Lower 1mm Stomata

epidermis/ which can be used on a local, regional or worldwide (Coleman frequancy in et al., 2010). DELTA is a modern software package that manages taxonomic research (Dallwitz, 2009). In this work DELTA is selected because it promotes consistency in descriptive taxon-

upper omy, can produce descriptive taxonomy in natural language epidermis %Stomatal

index of the ready for publication and produce conventional (dichotomous) keys. 2 upper 1mm Stomata epidermis/ frequancy in ) 2 Table 4 List of 8 characters and their character states recorded comparatively for studied 12 lower Area of species of the genius Amaranthus in the format of the DELLTA programs. Character epidermal cells (mm number is preceded by # and states of the same character are assigned serial . Leaf

) numbers. 2

#1. Types of stomata at the upper epidermis/

upper 1. anomocytic only/ Area of Amaranthus epidermal

cells (mm 2. anomocytic and hemiparacytic/ #2. Trichomes type/ 1. multicellular, uniseriate only, glandular and unbranched/

m) 2. multicellular, uniseriate and biseriate, glandular and unbranched/ l ( Lower thickness epidermis #3. Phloem strand in midrib/ 1. present/ 2. absent/ m)

l #4. Mesophyll organization/ ( Upper

thickness 1. isolateral/ epidermis 2. dorsiventral/ #5. Petiole shape in cross section/ m)

l 1. crescent (with two small protuberances)/ ( length

Palaside 2. V-shaped (without protuberances)/ 3. cordate (with two small protuberances)/ #6. Shape of vascular bundles in petiole/ m) 1. crescent/ l ( 2. V-shaped/ thickness Mesophyll #7. Bundle sheath in petiole/ 1. present/

m) 2. absent/ l ( 90.2290.08 61.56 81.8 20.5 22.88 15.65 8.49 10.33 9.99 0.98 0.66 0.58 1.54 84 80 14.3 12.2 70 82 9.74 11.81 1.55 1.44 121.02 139.27 51.51 46.93 140.86 120.86 28.99 15.23 11.21 1.34 1.31 72 4 66 10.28 2.72 292.6 72.19 101.46 29.02 44.18 16.14 17.92 3.5 2.84 40 10 64 18.18 1.41 201.72 48.97 159.03 128.95 35.24 17.57 12.43 2.38 2.66 26 8.28 36 14.28 1.48 197.03 59.74 117.71128.98 86.1 98.83 22.02 27.71 17.58 14.07 13.44 10.49 2.433 1.52 1.12 1.56 92 64 20.35 12.03 98 58 16.2.7 13.87 1.17 2.67 177.41 122.86 62.52 47.16 142.95 103.6 22.73 15.31 13.61 1.62 1.35 76 10.5 90 2.04 3.54 230.7 99.07 162.13145.49 128.02122.47 99.39 34.25114.48 97.53 33.11 15.37 85.58 32.8 15.04 17.04 27.54 16.67 12.94 12.06 3.33 12.11 9.06 1.24 2.85 1.97 2.89 0.76 88 1.52 2.24 86 16.92 50 64 11.71 50 10.36 8.98 98 8.47 36 84 13.8 3.28 5.88 2.62 12.72 262.92 2.66 289.29 73.45 1.19 238.14 155.51 181.06 78.3 49.87 Lamina average

thickness #8. Number of vascular bundles in petiole/ 1. 8/ 2. 4/ 3. 13/

x 4. 6/ A. albus A. blitum A. caudatus A. deﬂexus A. dubius A. graecizans A. hybridus A. powellii A. retroﬂexus A. spinosus A. tricolor A. ozanonii 5. 12/

Characters 6. 11/ 1 2 3 4 6 5 7 8 9 10 11 12 No Species 7. 7/ Table 3 Mean values for 14 quantitative characters of the leaf and stem of the studied species of

Co Co Vb

Pa Scr 1 2 Scr 100µm 100µm

Co Co

Vb BS Scr Vb

3 Pa 4 Pa 100µm Scr 100µm

Figs. 1–4. (1) TS in petiole of Amaranthus caudatus reveals that petiole is cordate in shape at TS with two small protuberances, epidermis is uniseriate, vascular bundles (Vb) have V-shape, ground tissue consists of angular collenchyma (Co) tissue below the epidermis followed by thin parenchyma (Pa), sand crystals (Scr) are present while druses absent and the bundle sheath (BS) is absent. (2) TS in Amaranthus tricolor petiole reveals that the small protuberances and bundle sheath are absent. (3 & 4) TS at petiole of Amaranthus spinosus and Amaranthus deﬂexus respectively shows that petiole at TS has an arc shape, vascular bundles have an arc shape, bundle sheath is present and the two small protuberances are clearly present.

Co Vb

Phs

Pa 5 6 100µ 100µm

Co Co D

Vb Pa Lv Scr Sc 7 8 Vb Pa 100µ 100 D St NKM

BS BS

St 9 KM 10 100µm 100µm

Figs. 5–10. (5) TS at the midrib of Amaranthus dubius shows that midrib is rounded in shape in TS, epidermis is uniseriate, ground tissue consists of angular collenchyma (Co) Tissue below the epidermis followed by thin parenchyma (Pa), united vascular bundles (Vb) with arc shape and phloem strand (Phs) below the upper epidermis which magnified in (6). (7) TS at the midrib of Amaranthus dubius shows that vascular bundles (separated) with rounded shape. (8) TS oat the midrib of Amaranthus spinosus shows that midrib has an arc shape at TS, sand crystals (Scr) present in midrib and druses (D) present at lamina. (9) Magnified portion of the lamina of Amaranthus tricolor shows kranz mesophyll (KM) and non-kranz mesophyll (NKM), isolateral mesophyll organization. (10) Magnified portion of the lamina of Amaranthus caudatus shows that epidermis uniseriate, dorsiventral mesophyll organization and druses (D). Lv = Lateral vein.

Figs. 11–16. (11 & 12) Surface view of the upper epidermis of Amaranthus retroﬂexus shows anomocytic stomata (st) and biseriate trichome (Tr). (13 & 14) Surface view of the upper epidermis of Amaranthus hybridus shows anomocytic, hemiparacytic stomata and uniseriate trichome. (15 & 16) Magniﬁed portion of the midrib of Amaranthus dubius and Amaranthus coadetus shows sand crystals (Scr) and druses (D) respectively.

Figs. 17 and 18. (17) T.S. of internode of Amaranthus powellii shows the uniseriate epidermis followed by few layers of angular chollenchyma (Co) and thin parenchyma (Pa) then an anomalous cambial ring (Aca) which found to move outwards and produce collateral vascular bundles with thin parenchyma as conjunctive tissue inwards to form the vascular system. Sand crystals (Scr) wear found. (18) T.S. of internode of Amaranthus deﬂexus shows that it has the same structure of other species but characterized by ligniﬁed parenchyma (Lpa) conjunctive tissue.

A.albus A.hybridus

A.blitum A.graecizans

A.Caudatus A.retroﬂexus

A.deﬂexus A. spinosus

A.tricolor A.dubius

1mm A. graecizans A.xoxanonii

Plate 1. Diagrams of the nodes transections of the studied species where leaf traces which diverged to the leaf bas are indicated by blackened xylem areas.

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